Aves Translation and Pronunciation Guide

Ben Creisler

Alexornis Brodkorb 1976 "Alex's bird"

al-ek-SOR-nis (Alex(ander) + Gr. ornis "bird") (m) named to honor Alexander Wetmore (1886--1978), American paleornithologist, "who has described more species of fossil birds than any other author". (?) Enantiornithes Alexornithidae L. Cret. NA. (Mexico)

Ambiortus Kurotchin 1982 "uncertain origin"

am-bee-OR-tus (from Lat. ambiguus "vacillating, uncertain" + Lat. ortus "origin, beginning")* (m) alluding to its uncertain placement in avian evolution, with a complex mosaic of archaic traits (a third phalange on the major digit of the wing) and modern features (fused carpometacarpals and a sternal keel like that found in carinate birds). Kurotchin thought the form was possibly ancestral to modern carinate birds. Ornithurae Ambiortidae E. Cret. CAs. (Mongolia)

Anatalavis Olson & Parris 1987 "duck-winged bird"*

an-a-TAY-la-vis (Lat. anat- (anas) "duck" + Lat. ala "wing" + Lat. avis "bird") (f) named for the shape of the humerus, resembling somewhat that found in modern ducks; for "Telmatornis" rex Shufeldt 1915. Anatalavis was originally based on two humeri (Holotype: YPM 902 (Yale Peabody Museum)) found in the Hornerstown Formation in New Jersey (dated either to the latest Cretaceous or earliest Paleocene), distinguished from Telamatornis by their much shorter, more robust and curved shafts. The recent discovery of very similar humeri associated with a partial skeleton (missing hindlimbs) and skull from the Eocene of England provides much more information on the genus (Olson 1999, Smithsonian Contributions to Paleobiology 89:231-243). The second species (Anatalavis oxfordi) is much larger and shares a number of characteristics with the modern Australian Magpie goose (Anseranas), most notably a V-shaped furcula with a large, deep symphysis. Anatalavis has a very broad, rounded, thin duck-like bill, indicating it was filter feeder with jaw muscles specialized for straining food (unlike the modern Magie goose, which has a strong, deep, hooked bill for digging outplant material); the short, robust humerus (with proportions more that of a falcon) suggests it was a strong, rapid flyer; the shape of the pelvis resembles that of wading birds rather than that of swimming duck-like birds. How much the Late Cretaceous type species would have resembled the early Eocene species is not clear without more skeletal material, but the new find suggests that specialized types of modern birds had already evolved by the end of the Mesozoic. The exact date of the basal Hornerstown Formation is still debated, and it may represent early Paleocene rather than Late Cretaceous.

Type Species: Anatalavis rex (Schufeldt 1915): "king," to represent a species of Telmatornis larger than Marsh's Telmatornis priscus--Olson and Parris (1987) erected the new genus Anatalavis for Schfeldt's species. Their original classification placed it in the order Charadriiformes, family Graculavidae. However, the new Eocene material indicates it is the earliest representative of the waterfowl order Anseriformes, closely related to the modern Magpie goose in the family Anseranatidae. Additional Species: Anatalavis oxfordi [OKS-for-die] Olson 1999: for Andrew Oxford, the English fossil collector who found the specimen in the early Eocene (Ypresian) London Clay at Walton-on-the-Naze, Essex, England. Ornithurae Anseriformes Anseranatidae Anatalavinae Late Cretaceous (or Paleocene) NA, Eocene Eur. [revised 8/2000]

Angelinornis Kashin 1972 "Angelina's bird"

AHN-gel-i-NOR-nis (Angelina + Gr. ornis "bird") (m) named to honor Angelina Mikhaylovna Sudilovskaya (1903- ), Russian ornithologist; to replace preoccupied Plegadornis Wetmore. [= Ichthyornis]

Apatornis Marsh 1873 "deceptive (vertebra) bird"

ap-a-TOR-nis (Gr. apatao "deceive" + Gr. ornis "bird")* (m) referring to the biconcave shape of its vertebrae, making them "deceptively" like the vertebrae found in fish, in contrast to the vertebrae with saddle-shaped articulations on the centra that are found in modern birds; originally identified as "Ichthyornis" celer. (See additional comments at Ichthyornis.) Ornithurae Apatornithiformes Apatornithidae L. Cret. NA.

Archaeopteryx von Meyer 1861 "ancient wing"

AHR-kee-OP-ter-iks (Gr. arkhaios "ancient" + Gr. pteryx "wing, feather") (f) Von Meyer first mentioned a fossil feather from the Jurassic Solnhofen deposits in a letter published in August, 1861. He described the find as either a flight or tail quill preserved in rock typical of the lithographic slates, though he was not completely convinced that the specimen came from the Mesozoic, since no bird fossils earlier than the Tertiary were then known. He did not propose a name for the isolated feather. In a follow-up letter published in September, 1861, after personal investigations, he confirmed that the feather was a genuine fossil from Jurassic deposits at Solnhofen. He also cited news that a nearly complete skeleton of a feather-bearing animal (now known as the "London specimen") had recently been found in the same lithographic quarries. In a slightly ambiguously worded passage, he gave "the animal" the name Archaeopteryx lithographica (lith-o-GRAF-i-ka, for the lithographic slates), the usual assumption being that the "animal" he intended to name was the source of his feather rather than the newly found "feather-bearing animal" whose specimen he apparently had not seen yet in person. In a fuller description of the feather published some months later, he took a cautious approach to exactly what sort of animal was the source of his feather, and was typically reluctant to conclude that the animal was a true bird, being a skeptical opponent of Cuvier's theories concerning the "correlation of parts." He did propose, however, that the feather came from an animal similar to the still formally undescribed and unnamed "feather-bearing animal." His German colleague J. A. Wagner gave a first description of the "feather-bearing animal" in 1861, proposing the name Griphosaurus "enigma lizard," although he only had seen a drawing, and not the specimen itself. The modern usage of the name Archaeopteryx was established by Richard Owen in 1863, when the "London specimen" was purchased by the British Museum. Owen accepted Cuvier's "law of correlation," and had no hesitation about identifying the feathered animal as a true bird. He also accepted von Meyer's generic name Archaeopteryx for the skeletal specimen with feathers, but thought a more appropriate species name would be macrura "long-tail" [mak-ROOR-a] in as much as this feature was highly distinctive--he reasoned that von Meyer's original species A. lithographica, based on a single feather, might well have been a short-tailed form, just as both long-tailed and short-tailed pterosaurs were known from the lithographic slates. Confusion over the correct type specimen and type species name was settled by an official decision of the International Commission on Zoological Nomenclature in 1964, which made the "London specimen" the holotype and Archaeopteryx lithographica the valid name. Although von Meyer did not specify if he intended Archaeopteryx to mean "ancient feather" or "ancient wing," the interpretation "ancient wing" is preferable, based on the standard usage of pteryx as "wing" for bird names in zoological nomenclature (Apteryx, etc.). It also would be consistent with the modern scientific usage of Archaeopteryx as defined by Owen. Von Meyer, moreover, accepted Owen's usage of the name for the "London specimen" (in Evans, 1865), although he still refused to recognize the animal as a true bird. Wellnhoffer (1993) has described a second slightly younger species (Archaeopteryx bavarica ba-VAR-i-ka "Bavarian") from a seventh specimen discovered at Solnhofen in 1992, distinguished by an ossified sternum and interdental plates in the jaws. (Placed on the Official List of Generic Names in Zoology with Name No. 1433 by ICZN Opinion # 607 based on the British Museum holotype.) Archaeopterygiformes Archaeopterygidae L. Jur. Eur. [dino-bird]

Archaeornis Petronievics in Petronievics & Woodward 1917 "ancient bird"

AHR-kee-OR-nis (Gr. arkhaios "ancient" + Gr. ornis "bird") (m) name proposed for the "Berlin specimen," found at Eichstätt; described as representing a genus supposedly distinct from the London specimen of Archaeopteryx. Reseachers now think Archaeopteryx had indeterminate growth more typical of reptiles, rather than the combination of rapid juvenile growth with a fixed adult size characteristic of modern birds, and that the minor differences between the London and Berlin specimens do not merit taxonomic separation. [= Archaeopteryx]

Asiahesperornis Nessov & Prizemlin 1991 "Asian hesperornithid"

AY-zha-HES-pe-ROR-nis (Asia + Hesperornis) (m) named to indcate a large flightless marine hesperornithid from the Late Cretaceous of Kazakhstan, Central Asia, the first hesperornithid known from Asia. Hesperornithiformes ?Hesperornithidae L. Cret. CAs.

Apsaravis Norell & Clark 2001 "apsara bird"

up-suh-RAY-vis (Sanskrit apsara, a mythical winged being + Lat. avis "bird")* (f) named for the apsara (also called apsaras), female winged consorts in Buddhist and Hindu art. Apsaravis is a primitive ornithurine bird known from a well preserved uncrushed, partly articulated partial skeleton with a fragmentary skull (Holotype: IGM 100/1017 (Institute of Geology, Mongolia)) from the Late Cretaceous (?Campanian) Ukhaa Tolgod site in southern Mongolia, dating from around 80 million years. Apsaravis has a mosaic of primitive and advanced features. It is notable as "the most basal avialan with an extensor process...related to the insertion of the extensor carpi radialis muscle and the propatagial ligaments in Aves"-- features that connect movement of the hand to movement of the forearm, with a key role in transition from the upstroke to the downstroke in the flight of modern birds. Other characters such as at least 10 ankylosed sacral vertebrae and the shape of the pelvis also place it close to modern birds, with some relation to the ornithurine toothed bird Ichthyornis, though Apsaravis may lack teeth. It also shares a number of primitive plesiomorphies present in Enantiornithes and some theropod dinosaurs, such as a strong concavity on the posterodorsal coracoid. Apsaravis comes from a continental deposit, indicating that early ornithurine birds were not confined to marine or near-shore environments as some earlier evidence suggested.

Type Species: Apsaravis ukhaana [oo-KAH-nuh] Norell & Clark 2001: (Ukhaa + Lat. - ana "belonging to") "from Ukhaa Tolgod," the type locality in southern Mongolia. Ornithurae Late Cretaceous (?Campanian) Mongolia [added 3-2001]

Avialae Gauthier 1986 "bird wings"

ay-vee-AY-lee (Lat. avis "bird" + Lat. ala "wing") (f) originally proposed by Gauthier for the clade composed of Archaeopteryx plus ornithurine birds, with the term Aves reserved for modern living birds and all the descendants of their most recent common ancestor. Padian (1997) revised Avialae to "define the stem group consisting of Neornithes [modern toothless birds] and maniraptorans closer to them than to Deinonychus [Deinonychosauria]," with the term Aves restored to a more traditional usage as a group that includes Archaeopteryx, plus extinct birds and all descendants of their most recent common ancestor. Other authors have treated Avialae as defined by Gauthier as a synonym of Aves. [clade]

Avisaurus Brett-Surman & Paul 1985 "bird lizard"

AY-vi-SAWR-us (Lat. avis "bird" + Gr. sauros "lizard") (m) named originally to indicate a supposed small dinosaur, with birdlike coossified metatarsals; reidentified as an enantiornithine bird by Bonaparte (1986). Enantiornithes Avisauridae L. Cret. NA.

Baptornis Marsh 1877 "diving bird"

bap-TOR-nis (Gr. bapto "dive" + Gr. ornis "bird") (m) named to indicate a flightless, swimming bird. Ornithurae Hesperornithiformes Baptornithidae L. Cret. NA.

Boluochia Zhou 1995 "for Boluochi (China)"

bwaw-lwaw-CHUHR-a (Boluochi + -a) (f) named for the town of Boluochi [bwaw-lwaw-chuhr], near where the incomplete holotype was found in Chaoyang County, Liaoning Province, Manchuria, northeastern China; notable for a hooked bill resembling that of modern predatory birds, a lack of teeth in the premaxillary, and strongly curved, sharply pointed raptorial claws. Enantiornithes i.s. E. Cret. China

Canadaga Hou 1999 "Canadian bird"*

KAN-uh-DAY-guh (Canada + ?Lat. - ago "having characteristics of" + -a) (f) named to indicate a fossil bird from northern Canada. Canadaga is a very large hesperornithid known from three neck vertebrae (Holotype: NMC 41050 (Canadian National Museum)), as well as referred material apparently belonging to juveniles (a caudal vertebra and parts of two femurs). The fossil material comes from Bylot Island, Northwest Territories, Arctic Canada, and dates from the Late Cretaceous (middle Maastichtian), thus representing the latest occurrence of a hesperornithid so far discovered. The preserved neck vertebrae likely are vertebrae 15-17; vertebra 16 has a centrum 28 mm long. The vertebrae show many similarities to those of Hesperornis but differ in a number of details, including having a fan-shaped expansion on the end of the centrum while the middle part of the centrum is constricted, and having a large and deep concavitas lateralis occupying almost the entire lateral surface of the centrum. The transverse process on the vertebrae has a deep cavity probably connected with the neural canal, a character which distinguishes Canadaga from other known hesperornithids. An adult Canadaga may have been more than 1.5 m (5 ft) long. The discovery of juvenile bones suggests suggests Canadaga may have bred in colonies in the high arctic, as suggested for some species of the earlier Hesperornis.

Type Species: Canadaga arctica [ARK-ti-kuh] Hou 1999: "arctic," referring to its discovery in the Arctic region of northern Canada, one of the most northerly occurrences of hesperornithids known. Ornithurae Hesperornithiformes Hesperornithidae Late Cretaceous (mid-Maastrichtian) NA. [added 7/2000]

Cathayornis Zhou, Jin & Zhang 1992 "Cathay bird"

KATH-ay-OR-nis (Cathay, old poetic name for China + Gr. ornis "bird") (m) named for Cathay, an old name for China; for a form more advanced than Archaeopteryx, found in the Jiufotang Formation, Chaoyang, Lianoning Province, China. Enantiornithes Cathayornithidae E. Cret. China

Caudipteryx Ji Q., Currie, Norell & Ji S. 1998 "tail feather"

kaw-DIP-ter-iks or kaw-dip-TAYR-iks (also KAW-dee-tayr-iks, Curries' own pronunciation) (Lat. cauda "tail" + Greek pteryx "wing, feather") (f) named to indicate a turkey-sized (3-ft.long) theropod notable for large symmetrical feathers arranged fanlike on the end of the tail. Caudipteryx is thought to represent a type of feathered ground-living dinosaur that may be related to the Oviraptorosauria. The two known specimens of the genus were found in the Jiulongsong Member of the Chaomidianzi Formation, Sihetun area of Liaoning Province, northeastern China, in ancient lake deposits that preserved impressions of rather large symmetrical shafted feathers on the tail and forelimbs, as well as downlike feathers on the body. However, the relatively small size of the forelimbs, the proportionately large hindlimbs and the symmetrical design of the feathers indicate the animal very likely could not fly. The animal is identified as a dinosaur rather than a bird based on a list of distinctive features, but most importantly, a skull with a bony bar behind the eyes (not found in birds) and other theropod characteristics. The pelvis is theropodlike (similar to dromaeosaurs and oviraptorids): the ischium lacks the posterodorsal process found in Archaeopteryx and other birds. Additionally, the hallux (first digit on the foot) is reduced and positioned high on the metatarsals, off the ground and oriented outward and slightly forward, as in typical theropod dinosaurs; by constrast, the hallux in most birds, including Archaeopteryx, is well-developed, low on the metatarsals and positioned backwards for perching or contact with the ground.

The two specimens were originally assigned to Protarcheopteryx. However, Caudipteryx differs from Protarchaeopteryx: its jaws are toothless except for usual hooked spike-like teeth at the tip of the upper jaw (premaxillary) that were probably partly covered with a horny beak; its tail is relatively short (one-quarter the length of the body, with 22 vertebrae (as in Archaeopteryx)); the forelimbs are relatively long for a nonavian theropod but shorter than in Protarchaeopteryx, with large secondary feathers attached to the second digit on the hand; it has a gizzard containing small grinding stones, indicating it may have eaten plant material.

Recent studies suggest Caudipteryx may be related to oviraptorosaurs. A few researchers have challenged the identification of Caudipteryx as a theropod dinosaur and propose that it is a secondarily flightless plant-eating bird that evolved from primitive Archaeopteryx-like forms. Type species: Caudipteryx zoui [DZOH-ie] "Zou's tail-feather" to honor Zou Jaihua, vice-premier of China, for his support of scientific work in Liaoning. Theropoda Maniraptora E. Cret. China [revised 6/99]

Ceramornis Brodkorb 1963 "Cretaceous bird"

SER-a-MOR-nis (Gr. keramos "potter's clay" + Gr. ornis "bird")* (m) based on the name Cimolopteryx (from Gr. kimolia "white chalky clay or earth," used with the applied meaning "Cretaceous") to indicate a closely related form. Ornithurae Charadriiformes Cimolopterygidae L. Cret. NA.

Changchengornis Ji Q., Chiappe & Ji S. 1999 "Great Wall bird"

chahng-chuhng-OR-nis (Chin. Chang Cheng (Great Wall) + Gr. ornis "bird")* (m) referring to the Great Wall of China (Chang Cheng), famous manmade structure erected south of the northeastern region of China in which the fossil was found (Liaoning Province, Sihetun-Jianshangou area); for a blue jay-sized, toothless confuciusornithid bird, differing from Confuciusornis in having a strongly curved beak and a lower jaw that is very deep at the back and much shorter than the skull; differing also in proportions of the metacarpals and hallux, and in the lack of an opening (foramen) on the deltopectoral muscle crest on the humerus. The large sternum is flat without a keel, overlapping the first two rows of a set of gastralia. The type specimen (GMV2129-a/b) in the National Geological Museum of China (Beijing) consists of two counterslabs of a nearly complete specimen preserving most of the plumage as a carbonized halo around the fossil bones. There is no tailfan, but the tail bears two long ribbon-like feathers, a feature also preserved in some specimens of Confuciusornis and possibly associated with sexual dimorphism (indicating males?), though it might have been found in both sexes and only missing in some Confuciusornis specimens because of immaturity or moulting.

The foot structure in Changchengornis indicates well-developed perching ability. Hou, Martin and Zhou (1996) have suggested that Confuciusornis (and presumably the closely related new genus) might have been specialized for climbing trees because of a proposed upright, squirrel-like posture and forelimbs with long-clawed digits. However, other researchers have disputed such an unusual posture (unlike any known bird or dinosaur), as well as the idea that the large curved digits on the wings were designed for climbing.

Type Species: Changchengornis hengdaoziensis [HUHNG-dowd-zuh-EN-sis] Ji Q., Chiappe, Ji S. 1999: for the Hengdaozi stratigraphic horizon (Early Cretaceous?) of the Chaomidianzi Formation. Metornithes Confusiusornithidae Late Jurassic-Early Cretaceous China [entry added 4-99]

Chaoyangia Hou & Zhang 1993 "for Chaoyang (China)"

chow-YAHNG-ee-a (Chaoyang + -ia) (f) named for Chaoyang, where the fossil was found in Liaoning Province, China; notable for large ossified uncinate processes on its ribs, otherwise unknown in non-ornithurine birds. i.s. E. Cret. China

Cimolopteryx Marsh 1889 "Cretaceous wing"

sim-oh-LOP-ter-iks (Gr. kimolia "white chalky earth [Cretaceous]" + Gr. pteryx "wing")* (f) named to indicate a fossil bird found in the Late Cretaceous deposits of Wyoming. Ornithurae Charadriiformes Cimolopterygidae L. Cret. NA.

Colonosaurus Marsh 1872 "stunted lizard"

ko-LON-o-SAWR-us (from Gr. kylos "stunted" + Gr. sauros "lizard")* (m) named proposed for a small toothed lower jaw found with the type specimen of Ichthyornis, and which Marsh first identified as belonging to a reptile; he later correctly attributed the dentary to Ichthyornis. [= Ichthyornis]

Concornis Sanz & Buscalioni 1992 "Cuenca Province (Spain) bird"

kong-KOR-nis (Conca, Latin name for Cuena Province, Spain + Gr. ornis "bird") (m) named to indicate a fossil bird found at Las Hoyas, Cuenca Province, Spain. Enantiornithes i.s. E. Cret. Eur.

Confuciusornis Hou, Zhou, Gu & Zhang 1995 "Confucius' bird"

kon-FYOO-shi-SOR-nis (Confusius + Gr. ornis "bird") (m) named to honor the great Chinese moral philosopher Confucius (551-479 B.P.); for an early form with a toothless beak that apparently evolved independently of modern birds. Known from dozens of newly discovered specimens, yet to be described. Enantiornithes i.s. E. Cret. China

Coniornis Marsh 1893 "Cretaceous bird"

kon-ee-OR-nis (Gr. konis "dust, chalk [Cretaceous]" + Gr. ornis "bird") (m) alluding to the Late Cretaceous (Judith River) deposits at Dog Creek, Montana, where the fossil was found, indicating that hesperornithids lived in freshwater as well as marine environments. Ornithurae Hesperornithiformes Hesperornithidae L. Cret. NA. [= ?Hesperornis]

Cuspirostrisornis Hou 1997 "pointed-rostrum bird"

KUH-spi-ros-tri-SOR-nis (Lat. cuspirostris "having a pointed rostrum" + Gr. ornis "bird") (m) referring to the long and slender rostrum on an enantiornithine toothed bird about the size of large sparrow; known from a complete, mostly articulated skeleton with skull (Holotype: IVPP V. 10897) found in the Jiufotang Formation at Boluochi, Chaoyang County, Liaoning Province, northeastern China. Skull 2.7 cm long, 5 teeth on each side of the premaxillary and tip of lower jaws; lower jaw slender and straight; humerus 2.9 cm long, with large internal condyle; ulna 3.2 cm long; radius 2.95 long; sternum has manubium and relatively developed keel; femur 2.73 cm long with large internal condyle; tibiotarsus 3.25 cm long; tarsometatarsus 1.9 cm long; toe-claws large and recurved.

Type Species: Cuspirostrisornis houi [HOH-ie] Hou 1997: for Hou Jifeng, senior engineer with the Institute of Vertebrate Paleontology and Paleoanthropology (Beijing). Enantiornithes Cuspirostrisornithidae Early Cretaceous (?Barremian) China [2/2000]

Enaliornis Seeley 1876 "sea bird"

e-NAL-ee-OR-nis (Gr. enalios "belonging to the sea" + Gr. ornis "bird") (m) proposed in 1869 to replace preoccupied Pelagornis Seeley 1866 "sea bird" (a nomen nudum), with a similar meaning; for an apparently flightless sea bird known from fragmentary remains that prove, according to Seeley (1885): "that some of the vertebrae are bi-concave in the back, that the avian form of articulation exists in the neck, and that the head resembles the divers, that the tibia is like that of the grebes, while the pelvis has some characters in common with the penguins." Ornithurae (?)Hesperornithiformes Enaliornithidae E. Cret. Eur.

Enantiornis Walker 1981 "opposite bird"

e-NAN-tee-OR-nis (Gr. enantios "opposite" + Gr. ornis "bird") (m) named for unique anatomical features not previously found in other known birds, in particular the articulation between the scapula and coracoid, where the "normal" configuration is completely reversed. Enantiornithes Enantiornithidae L. Cret. SA.

Enantiornithes Walker 1981 "opposite birds"

e-NAN-tee-or-NITH-eez (Gr. enantios "opposite" + Gr. ornith- (ornis) "bird" + -es) (m) Walker originally proposed the group Enantiornithes for the genus Enantiornis and some then unnamed specimens from Argentina, based on a suite of unusual anatomical features. Recent discoveries and new research have established that the Enantiornithes were a widespread and surprisingly diverse group of birds (ranging from sparrow- to turkey-vulture size, and including toothless forms) that flourished during the Cretaceous, but went extinct before the Tertiary along with the dinosaurs. A number of features distinguish so-called "opposite birds" from modern forms (Neornithes): "opposite birds" share toothed jaws and a rather primitive pelvic region with Archaeopteryx, but have a long pygostyle with a shortened, fused tail; the three tarsal elements in the feet have the opposite developmental fusion (proximodistal: "inward-out" relative to the midline of the body) from that found in modern birds (distal to proximal: "outward-in"); the shape of the triosseal canal (for the ligament responsible for the wing's upstroke) is distinctive; the scapula, coracoid and humerus, as well as some elements of the hind limbs also possess other unique features. Recent studies (Chinsamy, Chiappe, Dodson, 1994) of leg bones show the presence of growth rings, indicating that enantiornithines had a growth pattern, and perhaps a metabolism, that was slower or otherwise different from that found in modern fully endothermic birds, which have rapid juvenile growth and a fixed adult size. Flying forms of enantiornithes were nonetheless capable of sustained, powered flight, and had toes adapted to perching in trees. [taxon]/[clade]

Eocathayornis Zhou 2002 "dawn China bird"

EE-oh-kath-ay-OR-nis (Gr. eos "dawn" + Cathay "China" + Gr. ornis "bird") (m) named to indicate a bird from China more primitive than Cathayornis (the type specimen found in 1994 was originally identified as a fossil of Cathayornis). Eocathayornis is a small enantiornithine bird (about the size of a sparrow) known from bone impressions of a partial skeleton (disarticulated skull; neck, and upper part of the body, including complete forelimbs, coracoids, scapulae, sternum, ribs; 4 caudal vertebrae and part of a pygostyle) (Holotype: V10916 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), found in the Early Cretaceous (Barremian?) Jiufotang Formation, at Boluochi, in Chaoyang County, Liaoning Province, northeast China. The jaws have teeth; the neck vertebrae (probably 11) are heterocoelous; the sternum has a low keel and a pair of caudo-lateral processes (projecting backwards). The digits on the hand have 2 phalanges with a small claw on the short first finger (alular), 3 phalanges with a claw on the long second (main wing) finger and 2 phalanges with a small claw on the third finger; the manus (25.5 mm) is slightly shorter than the forearm, and the ulna (26 mm) is 110% the length of the humerus (23.5 mm). The advanced features of the scapula and wing suggest Eocathayornis had powerful flapping capability; it probably had an alula, found in a number of other early enantiornithine birds, for control in landing.

EocathayornisTypes Species: Eocathayornis walkeri [WAW-ker-ie] Zhou 2002: for the British paleontologist Cyril Alexander Walker, "who first published and recognized the significance of the Enantiornithes." Enantiornithes Early Cretaceous (Barremian?) China [added 6-2002]

Eoalulavis Sanz, Chiappe, Perez-Moreno, Buscalioni, Moratalla, Ortega & Poyato-Ariza 1996 "dawn alula bird"

ee-oh-AL-yuh-LAY-vis (or ee-oh-ay-LUHL-a-vis) (Gr. eos "dawn" + New Lat. alula "little wing [bastard wing]" + Lat. avis "bird") (f) referring to the early development of the alula or "bastard wing"--feathers attached to a movable "thumb" on the leading edge of the wing, used to smooth out the air-flow over the main wing, thus allowing better control in slow-speed flight (a feature apparently not present in Archaeopteryx); for a headless specimen about the size of a goldfinch, with well preserved feathers, and stomach contents. Enantiornithes E. Cret. Eur. (Spain)

Eoenantiornis Hou, Martin, Zhou & Fedducia 1999 "dawn opposite bird"

ee-oh-e-NAN-tee-OR-nis (Gr. eos "dawn" + enantios "opposite" + Gr. ornis "bird") (m) named to indicate one of the oldest known enantiornithine birds, intermediate in form between Archaeopteryx and Cathayornis. Eoenantiornis is based on a nearly complete skeleton about the size of a robin (Holotype: IVPP V11537 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), from the Yixian Formation (Early Cretaceous), Shihetun, Beipiao, Liaoning Province, China. The skull is short and deep. The small teeth are set in a groove (as in Hesperornis) and have short, unserrated crowns with waisted bases and expanded roots, the tooth form typical of other Mesozoic toothed birds. The neck is relatively long (11 vertebrae) compared to Archaeopteryx and Confuciusornis. The synsacrum is fused with 6-8 vertebrae; the pygostyle is long and tapering. The coracoid, sternum, and hand are intermediate in form between Archaeopteryx and enantiornithine birds; the outer digit has only one phalange. The humerus is about 95% the length of the ulna; the carpometacarpi are short (44% of the ulna). An alula is present, the earliest record of this feature used in avian flight control (also found in Eoalulavis from Spain).

Type Species: Eoenantiornis buhleri [BOO-le-rie] Hou, Martin, Zhou & Fedducia 1999; for the late Paul Buhler, "a distinguished German functional morphologist and paleornithologist." Enantiornithes Eoanantiornithidae Early Cretaceous China (added 7/99)

Eurolimnornis Kessler & Jurczak 1986 "European marsh bird"

YOOR-o-lim-NOR-nis (N. Lat. euro- "European" + Limnornis (Gr. limne "marsh, swamp" + Gr. ornis "bird")) (m) proposed for "a water bird...with flying capacity" found in Romania; redefined from fragments of humerus, ulna and carpometacarpus, material originally described as Limnornis Kessler & Jurczak 1984 (a preoccupied name), but minus the femur, which is now the type of Palaeocursornis. Limnorithiformes Eurolimnorthidae E. Cret. Eur. (Romania)

Gallornis Lambrecht 1931 "French bird"

ga-LOR-nis (Lat. Gallus, ancient inhabitant of France + Gr. ornis "bird") (m) referring to France, where the fossil was found at Auxerre; supposedly the oldest anatid [duck family], a disputed identification. Ornithurae ?Anatidae L. Cret. Eur. [nomen dubium]

Gansus Hou & Liu 1984 "Gansu (bird)"

GAHN-soos (Gansu + -us) (m) named for Gansu Province, China, where the fossil was found; among the earliest known remains of an apparent ornithurine bird. Ornithurae Gansuiformes Gansuidae E. Cret. China

Gargantuavis Buffetaut & Le Loeuff 1998 "Gargantua bird"

gar-gan-choo-AY-vis (t.L.m. gar-gahn-TOO-a-vis) (Gargantua, an imaginary giant in French medieval folk literature + Lat. avis "bird") (f) named after the imaginary giant Gargantua to indicate a giant flightless bird found in France; based on a broad, heavy 20 cm. x 18 cm. pelvis forming a sacrum with ten fused vertebrae, and a stout right femur about 15 cm. around, dated to 72 million years ago (Maastrichtian). The living animal would have been the size of a modern ostrich and would have weighed about 141 kg. (310 lbs.), making it the largest known bird from the Mesozoic. Because no remains of similarly large Cretaceous birds have been found in North America or Asia, it is possible that Gargantuavis evolved in isolation when western Europe was a large island during the Late Cretaceous. The discovery of a such a big bird is particularly interesting, since Gargantuavis may have laid some of the very large round fossil eggs found in southern France that scientists previously attributed to dinosaurs. Type species: Gargantuavis philoinos [fi-LOY-nos] "fond of wine" for the vineyards in the Languedoc region of southern France where the fossils were found. Aves L. Cret. Eur.

Gobipteryx Elzanowski 1974 "Gobi wing"

goh-BEEP-te-riks (Gobi + Gr. pteryx "wing") (f) named for the Gobi Desert, Mongolia, where the fossils were found (Kermeen Tsav); a toothless flying bird, known from eggs and embryos. Enantiornithes Gobipterygidae L. Cret. Mongolia

Graculavus Marsh 1872 "cormorant ancestor"

gra-KUHL-a-vus (Lat. graculus "cormorant" + Lat. avus "grandfather, ancestor")* (m) named to indicate a form "evidently most nearly allied to Cormorants" according to Marsh. Ornithurae Charadriiformes Graculavidae L. Cret. NA.

Griphosaurus Wagner 1861 "engima lizard"

GRIF-o-SAWR-us (for Gr. gryphos "enigma" + Gr. sauros "lizard")* (m) named to indicate its puzzling nature. Wagner says: "the identity of these epidermic structures with true birds' feathers is by no means proved; they might still only be peculiar adornments...I do not hesitate to regard this as a reptile of the order Sauria... Darwin and his adherents will probably employ the new discovery as an exceedingly welcome occurrence for the justification of the their strange views upon the transformation of animals....their views must at once be rejected as fantastic dreams with which the exact investigation of nature has nothing to do." (See additional comments at Archaeopteryx.) (Placed on the Official Index of Rejected and Invalid Generic Names as an objective synonym of Archaeopteryx by ICZN Opinion # 607) [= Archaeopteryx]

Gurilynia Kurochkin 1999 "for Gurilyn Tsav (Mongolia)"

goor-i-LEE-nee-uh (Gurilyn + -ia) (f) named for the Gurilyn Tsav locality, South Gobi Aimak, Mongolia, where the fossils were found in the Nemegt Formation. Based on fragmentary material: proximal end of a right humerus (Holotype: PIN 4499-12), distal end of left a humerus (PIN 4499-13), and shoulder-end of a left coracoid (PIN 4499-14). Gurilynia is "well distinguished from all other Enantiornithes for which the processes of the shoulder bones and the dorsal head of the coracoid are known." Both branches of the head of the humerus are about the same size, and the coracoid has an acute top of the coracoidal process and a very thin dorsal portion of the shaft. Gurilynia indicates that comparatively large enantiornithes were present in Central Asia by the end of the Late Cretaceous, though the entire size of the bird is difficult to estimate based on fragments alone.

Type Species: Gurilynia nessovi [NES-o-vie] Kurochkin 1999: for Lev Alexandrovich Nessov (1947-1995), Russian paleontologist, "in recognition of his invaluable contribution to the knowledge of fossil birds from Eurasia." Enantiornithes Late Cretaceous Mongolia (added 7/99)

Halimornis Chiappe, Lamb, & Ericson 2002 "marine bird"

hal-i-MOR-nis (Gr. halimos "belonging to the sea" + Gr. ornis "bird") (m) named to indicate a fossil bird found in marine deposits. Halimornis is a moderate-size enantiornithine bird (probably about as big as a pigeon), known from fragmentary remains, including parts of a right humerus, part of a right femur, 3 vertebrae and a pygostyle (Holotype: D2K 035 (Discovery 2000, Birmingham), plus additional elements (part of a left scapula, vertebral centrum and a neural arch (UAMNH PV996.1.1 (Alabama Museum of Natural History, Tuscaloosa) thought to be parts of the same individual as the holotype), found in the Late Cretaceous (Campanian) Mooreville Chalk Formation, Greene County, Alabama. The humerus has a bicipital crest that approaches the level of the humeral head (more proximally located than in any other enantiornithine) and an inflated area that projects laterally in the distal end of the femur. Halimornis is one of the few enantiornithine bird discovered in marine deposits, in a spot that would have been around 50 km (31 miles) out to sea from the ancient Campanian shoreline of the eastern margin of the Western Interior Seaway. Enantiornithines may have been more common in marine ecosystems than previously realized.

Types Species: Halimornis thompsoni [TOMP-so-nie] Chiappe, Lamb, & Ericson 2002: for Mrs. W. Thompson, the landlord of the area in which the specimen was found, in recognition of her many years of support for fossil collecting on the property. More correctly, the species names should be spelled thompsonae because it honors a woman. Ornithothoraces Enantiornithes Euenantiornithes Late Cretaceous (Campanian) NA [added 6-2002]

Hargeria Lucas 1903 "for Harger"

har-GEER-ee-a (Harger + -ia) (f) named to honor Oskar Harger, Marsh's assistant, who prepared the specimen [= Hesperornis].

Hesperornis Marsh 1872 "western bird"

HES-pe-ROR-nis (Gr. hesperos "western" + Gr. ornis "bird") (m) named for the location of the type fossil site, west of the Mississippi River, the traditional definition of the American West; for specimens found near the Smoky Hill River, Kansas, on a Yale collecting expedition. Ornithurae Hesperonithiformes Hesperornithidae L. Cret. NA.

Horezmavis Nessov & Borkin 1983 "Khorezm (Uzbekistan) bird"

ho-REZ-ma-vis (Horezm + Lat. avis "bird") (f) named for the Khorezm oasis, near where the fossil was found in the Karakalpuk Autonomous Region, Uzbekistan. Enantiornithes Gobipterygidae E. Cret. Cas.

Iberomesornis Sanz & Bonaparte 1992 "Iberian intermediate bird"

ie-BER-o-me-SOR-nis (Lat. Iberia, name for the Iberian peninsula (modern Spain and Portugal) + Gr. mesos "middle, intermediate" + Gr. ornis "bird")* (m) named to indicate a form anatomically intermediate between Archaeopteryx and later birds, found at Las Hoyas, La Cierva Township, Cuenca Province, east-central Spain. Ornithothoraces Iberomesornithiformes Iberomesorthidae E. Cret. Eur. (Spain)

Ichthyornis Marsh 1872 "fish (vertebra) bird"

ik-thee-OR-nis (Gr. ikhthys "fish" + Gr. ornis "bird")* (m) As indicated by Marsh, the genus possessed "biconcave vertebrae, from which fish-like characteristic the name is derived." Its biconcave-shaped vertebrae contrasted with the saddle-shaped articulations on the centra of the vertebrae that are characteristic of modern birds, thus the Latin type species name I. dispar DIS-par "unlike". Marsh (Odontornithes (1880)) thought its biconcave vertebrae were of great evolutionary importance: "While all existing birds, and all of the extinct forms so far as known, including Hesperornis, have the peculiar saddle-shaped vertebrae, those of Ichthyornis, and its near ally Apatornis, are biconcave. This form is seen in a few recent and in many extinct Reptiles, and in the Amphibians; but is especially characteristic of Fishes, from which class it was undoubtedly inherited by the superior groups. This character alone indicates unmistakably a great antiquity for the class of birds." (He proposed a possible Palaeozoic origin for birds.) Marsh originally identified a toothed jaw found with the type material as that of a small reptile he called Colonosaurus "stunted lizard," but after further preparation of the specimen in 1873, decided the toothed jaw belonged to Ichthyornis itself. He later classified Hesperornis (found to have toothed jaws in 1875) along with Ichthyornis in the suborder Odontornithes "toothed birds." Controversy over whether Ichthyornis had teeth was revived in 1952 by J. T. Gregory, who reidentified Marsh's original toothed jaw as that of a juvenile mosasaur (Clidastes), and argued that such large toothed jaws seemed unlikely on an active flying bird. Gregory's position was widely accepted, and nearly all researchers reclassified Ichthyornis as a toothless bird, leading artists to revise the traditional restoration with a smaller tern-like beak. In 1967, however, M. V. Walker carefully restudied Marsh's original jaws, and concluded, that lacking further proof, the jaws should be considered to belong to Ichthyornis. Walker then found another partial skeleton with toothed jaws. An additional jaw specimen was identified among material attributed to Hesperornis (Gingrich 1972), and scholars no longer question the presence of teeth in Ichthyornis. (Ichthyornis does NOT mean "fishing bird" for its diet, as often stated.) Ornithurae Ichthyornithiformes Ichthyornithidae L. Cret. NA. (The supposed presence of Ichthyornis in Asia (Nessov) has been refuted (Koruchkin 1995).)

Jibeinia Hou 2000 (1997) "Jibei bird"*

jee-BAY-nee-uh (Chin. Jibei, a name for northern Hebei Province + -ia) (f) named to indicate a fossil bird from northern Hebei Province, northern China, the first Mesozoic bird found in Hebei Province. Jibeinia is known from a nearly complete skeleton (missing the back of the skull) preserved with an outline of feathers, found in a quarry (Dongtuyao) at Tuyao village, Luanjitu township, Fengning County, in northern Hebei Province, northern China in the Early Cretaceous Yixian Formation. The specimen represents a relatively primitive toothed bird about the size of a sparrow: skull 2.6 cm long, jaws provided with many pointed, unserrated teeth; ?11 cervical vertebrae (only 6 visible), with non-heterocoelous centra; sternum wide--xiphial (median) process long, lateral processes not expanded; humerus 2.33 cm long; radius 2.42 cm long; ulna 2.4 cm long; carpometacarpus bones not fused, but close to one another; 3rd finger with joints reduced; small claws on all 3 fingers; ?13 thoracic vertebrae; 8 sacral vertebrae; pubic bone tips fused but not expanded; femur 2.22 cm long; tibiotarsus 2.8 cm long; tarsometatarsus 1.63 cm long; metatarsal bones not completely fused at proximal ends, distal ends not fused; metatarsal 2 shorter than metatarsals 3 and 4; 10-12 caudal vertebrae, pygostyle present.

Hou classifies Jibeinia in the Sauriurae as a form more advanced than Confuciusornis, on a lineage leading to birds commonly classified in the Enantiornithes. The metatarsals show some fusion at the upper proximal ends, resembling the proximal fusion of the tarsometatarsals in the Enantiornithes. Jibeinia differs from Confuciusornis not only in having teeth, but in the construction of the hand and fingers, and the well developed sternum. The construction of the hand is much more like that of Sinornis than Confuciusornis--the finger bones are all distinct (no fused carpals) with small claws on each finger, including the reduced third digit. (Sinornis lacks a claw on the reduced third digit, while Confuciusornis has enlarged claws on the first and third digits, the joints of the third digit being long and relatively well developed.) The sternum in Jibeinia is more advanced than the simple sternum of Confuciusornis, which lacks the posterior lateral processes and the xiphial process found in Jibeinia.

Although a detailed description of "Jibeinia" was published in 1997 (Hou, L. 1997. Mesozoic Birds of China, pgs. 99-110), the original Chinese text did not provide the Latin name for the taxon, referring to the new fossil animal only in Chinese as the "Jibei niao [Jibei bird]." The Latin binomen "Jibeinia luanhera" only appeared in captions for illustrations, so the scientific name Jibeinia was technically not an available name under ICZN rules (See ICZN (4th Edition) Art. 13.6.1.). The Latin name Jibeinia luanhera appeared in the text of Hou's 2000 book Picture Book of Chinese Fossil Birds (pg. 54) with a citation of Hou's 1997 book, making the scientific name available as of 2000, although no catalog number is indicated for the holotype.

Type Species: Jibeinia luanhera [LWAHN-huh-ruh] Hou 2000 (1997): for the Luan River [Luan He] that runs from Inner Mongolia through Hebei Province and Jehol in northeastern China. Ornithothoraces i.s. Early Cretaceous (?Barremian) China [added 7/2000]

Judinornis Nessov & Borkin 1983 "Yudin's bird"

YOO-di-NOR-nis (Judin [= Yudin] + Gr. ornis "bird") (m) named to honor Konstantin Alekseyevich Yudin (1912-1980), noted Russian ornithologist and systematist. Ornithurae Hesperornithiformes Baptornithidae L. Cret. Mongolia

Jurapteryx Howgate 1985 "Jura wing"

ju-RAP-ter-iks (Jura + Gr. pteryx "wing, feather") (f) "from Jura, a common name for the Jurassic Alb where the specimen was discovered, and the name of the museum where the specimen is housed, (also indicative of the Jurassic age of the specimen)"; for the "Eichstätt specimen," described as a new genus from a small specimen. It is now thought that Archaeopteryx had a reptilian type of growth pattern, with gradual indeterminate growth, unlike modern birds, which have rapid early growth and determinate adult size. Most authorities therefore do not consider the variation in size among specimens attributed to Archaeopteryx to be a valid basis for making taxonomic distinctions. [= Archaeopteryx]

Kizylkumavis Nessov 1984 "Kizylkum (Kazakhstan) bird"

kiz-il-KOOM-a-vis (Kizylkum + Lat. avis "bird") (f) named for the Kizylkum Desert, Navoi District, Kazakhstan, where the fossil was found. ?Enantiornithes ?Alexornithidae L. Cret. CAs.

Kuszholia Nessov 1992 "birds' road"

koos-ZHOH-lee-a (Kazakh kus zholi "birds' road" [= Milky Way] + -ia) (f) named to a indicate a flightless or near flightless bird, or possibly a birdlike theropod, found at Dzhyrakuduk, Uzbekistan. The name derives from the Kazakh term for the Milky Way ("the birds' road"), and alludes to the birdlike nature of the animal. ?Aves Kuszholiidae L. Cret. (Uzbekistan)

Laornis Marsh 1870 "fossil bird"

lay-OR-nis (Gr. laos "stone" + Gr. ornis "bird") (m) for a tibiotarsus from Birmingham, New Jersey, belonging to a wading bird "nearly as large as a swan." Ornithurae Charadriiformes Graculavidae L. Cret. NA.

Largirostrornis Hou 1997 "large-rostrum bird"

LAHR-ji-ros-TROR-nis (Lat. largus "large" + Lat. rostrum + Gr. ornis "bird") (m) referring to the large and long rostrum on an enantiornithine toothed bird about the size of a large sparrow; known from a fairly complete, partially articulated skeleton with skull (Holotype: IVPP V. 10531) found in the Jiufotang Formation near Boluochi, in Chaoyang County, Liaoning Province, northeastern China. Skull is long (3.2 cm), with a braincase shorter than the large rostrum; teeth recurved, found only in the tip of the jaws, with 6 on each side of upper jaws; vertebrae amphicoelous with high spinal crest; sternum has long lateral process, expanded distally; furcula short and robust; coracoid long, arc-shaped articulation to sternum; humerus S-shaped, 3.1 cm long, with a capital groove; ulna 3.15 cm long; carpometacarpus mostly fused; claws on manual digits; fused sacrum; femur 2.85 cm long; tibiotarsus 3.3 cm long; tarsometatarsus 1.9 cm long; pygostyle present.

Type Species: Largirostrornis sexdentoris "six-toothed" Hou 1997: for the 6 teeth in each side of the upper jaw. Enantiornithes Cathayornithiformes Cuspirostrisornithidae Early Cretaceous (?Barremian) China [2/2000]

Lectavis Chiappe 1993 "Lecho (Formation) bird"

LEK-ta-vis (or lek-TAY-vis) (Lat. lectus "bed" (for Spanish lecho "bed") + Lat. avis "bird")* (f) named in reference to the Lecho Formation at El Brete in Salta Province, northwestern Argentina, where the fossil was found. Enantiornithes i.s. L. Cret. SA.

Lenesornis Kurochkin 1996 "Lev Nessov's bird"

LEN-e-SOR-nis (Le(v) + Nes(sov) + Gr. ornis "bird") (m) named to honor Lev Alexandrovich Nessov (1947-1995), Russian paleontologist, "in recognition of his contribution to the understanding of the paleontology of Eurasia"; for "Ichthyornis" maltshevskyi Nessov 1986, based on the front half of a synsacrum (PO 3434) from Dzhyrakuduk, Kizylkum Desert, Navois District, Bukhara Region, Uzbekistan; Bissektinskaya Suite (Coniacian). The preserved part of the synsacrum is wide for its entire length, unlike in Zyraornis, and has rib processes that form acute angles. The specimen shares "some derived characters with Nanantius," and thus belongs to the Enantiornithes "opposite birds" rather than to the Ichthyornithidae as Nessov (1986) originally proposed.

Type species: Lenesornis maltshevskyi [mal-CHEV-skee-ie] (Nessov 1986)

Enantiornithes Alexornithiformes Alexornithidae Late Cretaceous (Coniacian) CAs. [entry added 11-98]

Lestornis Marsh 1876 "robber bird"

les-TOR-nis (Gr. lestes "robber, thief" + Gr. ornis "bird") (m) [= Hesperornis]

Liaoningornis Hou 1997 "Liaoning Province (China) bird"

LYOW-ning-OR-nis (Liaoning (Chin. liao "distant" + Chin. ning "tranquil") + Gr. ornis "bird") (m) named to indicate a fossil bird found in an ancient lake deposit in Liaoning Province, northeastern China (one-time Manchuria). The sparrow-sized form is known from a partial skeleton with a well developed keeled breastbone for anchoring flight muscles, a short crow-like tarsometatarsus, and feet with a large hallux and sharp recurved claws designed for perching-- all advanced features that Archaeopteryx lacked. Its long and broad sternum, and ribs with uncinate processes are like those of modern birds, and suggest that Liaoningornis had an efficient air-sac system for breathing that would have permitted the high oxygen consumption needed for vigorous flight. It may well have had a true warm-blooded metabolism in contrast with both Archaeopteryx and enantiornithine birds such as Confuciusornis, which lacked air-sacs and show evidence of a reptilian-type of growth pattern. However, Liaoningornis also had primitive toothed jaws like many other Mesozoic birds. The date of the ancient lake deposit is crucial to the fossil bird's role in evolution. In an on-going controversy, different researchers have dated the Liaoning Province quarries either to the Late Jurassic (137 to 142 million years ago) or to the Early Cretaceous (121 million years ago). The Early Cretaceous date is supported by new radiometric dating--a Late Jurassic date would make the new form a near-contemporary of Archaeopteryx, indicating that modern ornithurine birds may have evolved much earlier than many experts now accept, thus relegating Archaeopteryx to an evolutionary dead end unrelated to modern forms. Liaoningornithiformes Liaoningornithidae (?L. Jur.) E. Cret. China

Liaoxiornis Hou & Chen 1999 "Liaoxi bird"

lyow-shee-OR-nis (Laioxi (Chin. liao "distant" + xi "west") + Gr. ornis) (m) named for Liaoxi, term for the western part of Liaoning Province, China, the region in which the fossil was found (Yixian Formation). Liaoxiornis is the smallest adult Mesozoic bird known (skeleton about 8.5 cm. (3.3 in.) long, the size of a large hummingbird). The holotype is a nearly complete skeleton (IVPP 130723 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), with some bones preserved as impressions. The skull is 24 mm long, tall and short with large orbits. The jaws have small teeth (5 in the lower jaw, 6 on the maxilla and premaxilla). The neck is long, with over 10 vertebrae. The sternum has a peculiar gingko-leaf shape not found in any other birds, with a low keel. The pubis lacks the pubic foot found in other primitive birds. The tail is long (25 mm), proportionately the longest bony tail known next to Archaeopteryx; the pygostyle is about 3/4 the length of the tail (18 mm), and the 8 sacral vertebrae are not fused. The forelimbs are relatively advanced, with fused carpals and metacarpals; the digits have a reduced number of phalanges and apparently lack claws. The structure of the hindlimbs is very primitive: the femur (16 mm) is longer than the humerus (15 mm), but only slightly shorter than the tibiotarsus (18 mm), which is simple and not strong. The feet have claws that are relatively long but not strongly curved. Liaoxiornis shows a mix of primitive and advanced features, resembling Archaeopteryx in some details, enantiornithes in others, and may prepresent an evolutionary link between Archaeopteryx and the enantiornithes. It was evidently a better flier than Archaeopteryx, with more advanced forelimbs and a shorter tail.

Type Species: Liaoxiornis delicatus [del-i-KAY-tuhs] Hou & Chen 1999 "delicate" for its small size. Sauriurae Early Cretaceous China (added 7/99)

Limenavis Clarke & Chiappe 2001 "threshold bird"

lie-men-AY-vis (or lie-MEN-a-vis) (Lat. limen "threshold" + Lat. avis "bird")* (f) named "for the window it offers into the origin of the radiation of the avian crown clade." Limenavis is a carinate bird based on various parts of a right wing (including end parts of humerus, ulna, carpometacarpus, digit II) (Holotype: PVL 4731 (Paleontologia de Vertebrados, Instituto Miguel Lillo, Tucuman)) found in the Late Cretaceous (Campanian-Maastrichtian) Allen Formation at Salitral Moreno, Rio Negro Province, south-central Argentina. Features such as a fossa on the dorsal, distal extremity of the humerus; distal fusion of metacarpals II and III; and an extensor process on metacarpal I, plus the carinate synapomorphies of a truncate contact of the dorsal trochlear surface of the ulna with the ulnar shaft and the loss of a tubercle adjacent to the tendinal groove on the distal ulna would indicate that Limenavis is closer to modern birds than Ichthyornis is, but is just outside the avian crown clade. Distinctive muscle attachment pits and ligament scars on the ulna, plus the location of the pisiform process, establish it as a new genus and species.

Type Species: Limenavis patagonica [pat-a-GOH-nik-uh] Clarke & Chiappe 2001 "from Patagonia" in Argentina, the region in which the specimens were found. Avialae Carinatae Late Cretaceous (Campanian-Maastrichtian) SA [added 8-2001]

Limnornis Kessler & Jurczak 1984 "marsh bird"

lim-NOR-nis (Gr. limne "marsh, swamp" + Gr. ornis "bird") (m) proposed for "a waterfowl species with developed wings, of the size of a larger grebe...The adaptation to the aquatic environment is indicated by the distal fragment of femur." However, the femur was later made the type of Palaeocursornis. (Preoccupied by Limnornis Gould 1839. See Eurolimnornis.)

Limosavis Shufeldt 1915 "Limosa bird"

li-MOH-sa-vis (Limosa, New Latin genus name for the godwit (from Lat. limosus "muddy") + Gr. avis "bird") (f) proposed replacement name for "inappropriate" Graculavus, since the fossil appeared to be related to godwits rather than cormorants. [= Graculavus]

Lingyuanornis Ji & Ji 1999 "Lingyuan bird"

ling-yooen-OR-nis (Chinese Lingyuan + Gr. ornis "bird") named for the city of Lingyuan, near where the specimen was found in Liaoning Province, China. Lingyuanornis is based on the counterpart to the holotype specimen of Liaoxiornis: the original specimen was split into mirror-image slab and counterslab, each half being acquired by a different institutions in Beijing (Liaoxiornis specimen by the Institute of Vertebrate Paleontology and Paleoanthropology and Lingyuanornis specimen by the National Geological Museum of China). Researchers then gave the specimens separate catalog numbers and described under them different scientific names. The name Liaoxiornis has priority by about a month: the original description of Liaoxiornis appeared in Chinese in the early February 1999 issue of Kexue Tongbao 44(3):311-315; the description of Lingyuanornis appeared in the March 1999 issue of Zhongguo Dizhi [Chinese Geology] 262:45-48. Note that the official English language description of Liaoxiornis did not appear until early May 1999 in Chinese Science Bulletin 44 (9):834-838, which made no mention of the earlier February publication date in Chinese in Kexue Tongbao.

Type Species: Lingyuanornis parvus [PAHR-vuhs] Ji & Ji 1999 "small" [= Liaoxiornis] [added 8-2001]

Lonchodytes Brodkorb 1963 "Lance diver"

long-KOD-i-teez (Gr. logkhos "lance" + Gr. dytes "diver")* (m) named for the Late Cretaceous Lance Formation, Niobrara County, Wyoming, where the fossil was found. Ornithurae Charadriiformes Cimolopterygidae L. Cret. NA.

Longchengornis Hou 1997 "Longcheng bird"

loong-chuhng-OR-nis (Chin. Longcheng "Dragon Town" + Gr. ornis "bird") (m) for Longcheng "Dragon Town," an ancient name for the modern city of Chaoyang (founded in 341 AD by Murong Kuang, a leader of the Xianbi (a group of Mongol people), as capital of the one-time Kingdom of Yan in modern Liaoning Province). Known from an incomplete skeleton with fragments of the skull (Holotype: IVPP V. 10530), from the Jiufotang Formation at Boluochi, Chaoyang County, Liaoning Province, northeastern China. A enantiornithine toothed bird about the size of a sparrow with a ventral crest on cervical vertebrae; humerus 3.25 cm long, with round depression on proximal end; radius 2.9 cm long; coracoid small, narrow and straight at distal end; furcula thin; vertebrae amphiplatyan; 8 sacrals; 15 caudal vertebrae, long, unfused; femur 2.15 cm long; tibiotarsus slender, 3.4 cm long; tarsometatarsus 2.15 cm long; pedal claws large, curved.

Type Species: Longchengornis sanyanensis [sahn-ye-NEN-sis] Hou 1997: "from Three Yan (states)" referring to three feudal states of Yan (Qin Yan, Hou Yan and Northern He Yan) that existed in the Chaoyang region during the Sixteen Kingdoms Period (316-452 AD) in northern China. Enantiornithes Cathayornithiformes Cathayornithidae Early Cretaceous (?Barremian) China [added 2/2000]

Longipteryx Zhang, Zhou, Hou & Gu 2000 "long wing"

lon-JIP-te-riks (Lat. longus "long" + Gr. pteryx "wing, feather") (f) named to indicate an enantiornithine bird with wings that are significantly longer than the hindlimbs (wings more than 1.5 times as long as back legs). Longipteryx is known from a complete articulated skeleton with feather impressions (Holotype: V 12325 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), as well as a second complete skeleton and additional bones, found in the Early Cretaceous (Barremian?) Jiufotang Formation near Chaoyang City in Liaoning Province, China. It roughly resembles a small kingfisher (around 20 cm (8 in) long from jaw-tip to end of pygosyle) in having powerful wings, relatively short hind limbs, a long head and an ability to perch. Its skull is about 6 cm (2.3 in) in length, with long jaws (premaxilla 70% of total length of skull; dentary 57% length of the skull) containing short, curved conical teeth. The neck had an estimated 9 vertebrae (7 preserved) mostly heterocoelous in form; the neck was relatively long and flexible, indicating that Longipteryx may have relied mainly on its head and neck in attacking prey. There are 10 thoracic vertebrae and at least 8 vertebrae in the synsacrum. The tail has 4 unfused vertebrae and a fused pygostyle longer than the total length of the free caudals. Gastralia are present. The thoracic ribs have unattached uncinate processes (the only enantiornithine found with uncinate processes), suggesting Longipteryx had a strong thorax and respiration cability similar to modern birds. Its big sternum with a prominent carina and its long wings also suggest it had stronger flying abilities than other primitive birds, although it also had claws on its 3 wing-fingers. Its long, opposable hallux suggest it spent time in trees when resting. Longipteryx differs from all other known enantiornithines in being specially adapted for feeding on aquatic prey and may indicate that enantiornithines originated before the Early Cretaceous to reach such early diversity. Note that the original description in Chinese was published in December, 2000 (Kexue Tongbao 45(24):2650-2657), the correct citation date; the paper appeared in English translation in June, 2001 (Chinese Science Bulletin 46(11):945-950).

Type Species: Longipteryx chaoyangensis [chow-yahng-EN-sis] Zhang, Zhou, Hou & Gu 2000 "from Chaoyang City," where the specimens were collected in Liaoning Province, northeast China. Enantiornithes Longipterygiformes Longipterygidae Early Cretaceous (Barremian?) China [added 8-2001]

Mononychus Perle, Norell, Chiappe & Clark 1993 "one claw"

mo-NON-i-kus (Gr. mono- "one, single" + Gr. onykh- (onyx) "claw" + -us) (m) named for a short, robust forelimb possessing a single stout claw "...unlike the delicate, recurved manus claws of most theropods." (Preoccupied by Mononychus Schueppel in Germar 1824, a conserved name under ICZN Opinion # 1529). [= Mononykus]

Mononykus Perle, Norell, Chiappe & Clark 1993 "one claw"

mo-NON-i-kus (arbitrary respelling of Mononychus (Gr. mono- "one, single" + Gr. onykh- (onyx) "claw" + -us) "one claw") (m) replacement name for preoccupied Mononychus Perle, Norell, Chiappe & Clark, named for a short, robust forelimb possessing a single stout claw "...unlike the delicate, recurved manus claws of most theropods." The function of such a strange modification of a wing or forelimb in conjunction with a running bipedal posture is unclear--the large claw may have been used for digging, or perhaps for tearing into bark, termite mounds or insect nests after food. The genus is possibly related to Alvarezsaurus from Argentina (Novas, 1994), a form generally classified as a dinosaur. The identification of Mononykus as a bird rather than a dinosaur is disputed by a number of researchers, but a newly discovered specimen with a complete skull appears to support bird status--the orbit is connected to the infratemporal fenestra, a condition known only for birds among archosaurs. Metornithes L. Cret. Mongolia [dino-bird]

Nanantius Molnar 1986 "dwarf opposite (bird)"

na-NAN-tee-us (Gr. nanos "dwarf" + from Gr. (en)antios "opposite")* (m) referring to its small size (blackbird- or American robin-size) compared to Enantiornis (turkey vulture-size) and some related forms, and to its classification in the Enantiornithes "opposite birds." Enantiornithidae E. Cret. Aus.

Neogaeornis Lambrecht 1929 "New World bird"

NEE-o-jee-OR-nis (Gr. neos "new" + Gr. gaia "earth" + Gr. ornis "bird") (m) named to indicate the first Cretaceous bird discovered in the Southern Hemisphere (found in Chile). Now recognized as the earliest known loon (Olson 1992). Ornithurae Graviidae L. Cret. SA.

Neuquenornis Chiappe 1994 in Chiappe & Calvo 1994 "Neuquén (Argentina) bird"

nayoo-ke-NOR-nis (Neuquén + Gr. ornis "bird") (m) named for the city of Neuquén, Neuquén Province, Argentina, where the holotype specimen was found in the Late Cretaceous Rio Colorado Formation. The Latin type species name volans VOL-anz "flying" refers to its inferred flying capability. Enantiornithes Avisauridae L. Cret. AS.

Noguerornis Lacasa Ruiz 1989 "Noguera River (Spain) bird"

noh-ge-ROR-nis (Noguera + Gr. ornis "bird") (m) named for the Riu Noguera (River), near Montsech, Lerida Province, Catalonia, Spain, where the fossil was found; more adapted to powered flight than Archaeopteryx, with elements of the manus interlocking in a rigid structure, distal portions of the wing elongated, and the wing surface widened. i.s. E. Cret. Eur. (Spain)

Ornithothoraces Chiappe & Calvo 1994 "bird chests"

or-NITH-o-thoh-RAY-seez (Gr. ornith- (ornis) + Gr. thorak- (thorax) + -es) (m) referring to the modifications of the shoulder girdle and thorax for powered flight; including Iberomesornis, Neornithes and all descendents of their common ancestor. [clade]

Ornithurae Haeckel 1866 "Bird tails"

or-ni-THOOR-ee (Gr. ornith- (ornis) "bird" + Gr. oura "tail") (f) Proposed for birds with short tails and a pygostyle, distinct from the Saururae (Archaeopteryx) with long reptilian type tails. Redefined by Gauthier 1986 as a clade including "extant birds and all other taxa, such as Ichthyornis and Hesperornis, that are closer to extant birds than is Archaeopteryx." [taxon]/[clade]

Otogornis Hou 1994 "Otog-qi (Inner Mongolia) bird"

OT-o-GOR-nis (Otog + Gr. ornis "bird") (m) named for the fossil locality: Otog-qi, Yikezhao-meng, Inner Mongolia, China; to date, the earliest record of avian fossils from China. ?Enantiornithes E. Cret. CAs. (Inner Mongolia)

Palaeocursornis Kessler & Jurczak 1986 "ancient running bird"

PAY-lee-o-kur-SOR-nis (Gr. palaios "ancient" + Lat. cursor "runner" + Gr. ornis "bird") (m) proposed for a "running bird form of smaller size...the earliest known remains of the Ratitae," based on a femur fragment originally included in Limnornis Kessler & Jurczak, but later recognized as resembling femora from subfossil and living species of Ratite birds. Palaeocursornithiformes Cursornithidae E. Cret. Eur (Romania) [nomen dubium]

Palaeopteryx Jensen 1981 "ancient wing"

PAY-lee-OP-ter-iks (Gr. palaios "ancient" + Gr. pteryx "wing") (f) name proposed for a "tibiotarsus" belonging to a supposed bird. Jensen and Padian (1989) reidentified the specimen as the distal radius of an indeterminate maniraptoran theropod, and tried to declare the name invalid on the grounds that the taxon had not been properly diagnosed when proposed. Theropoda Maniraptora L. Jur. NA. [nomen dubium] [dino-bird]

Palaeotringa Marsh 1870 "ancient shore bird"

PAY-lee-o-TRING-ga (Gr. palaios "ancient" + Gr. tryggas "a shore bird" (mentioned by Artistotle))* (f) named to indicate a Late Cretaceous wading bird "apparently belonging to the Grallae [obsolete name for wading birds]." Ornithurae Charadriiformes Graculavidae L. Cret. NA.

Palintropus Brodkorb 1970 "backward-bender"

pa-LIN-tro-pus (Gr. palin "backwards" + Gr. tropo "bend" + -us) (m) named "in reference to the recurved facet on the coracoid"; for "Cimolopteryx" retusus Marsh; possibly related to Apatornis. Ornithurae Charadriiformes Palintropidae L. Cret. NA.

Parahesperornis Martin 1984 "near-Hesperornis"

PAR-a-HES-pe-ROR-nis (Gr. para "near" + Hesperornis (Gr. hesperos "western" + Gr. ornis "bird"))* (m) named to indicate a foot-propelled diving bird contemporary with Hesperornis; for a specimen previously identified as "Hesperornis gracilis." Ornithurae Hesperornithiformes Hesperornithidae L. Cret. NA.

Parascaniornis Lambrecht 1933 "near-Scaniornis"

PAR-a-SKAN-ee-OR-nis (Gr. para "near" + Scania, Latin name for the Scandinavian region + Gr. ornis "bird") (m) named for its resemblance to the Paleocene Scaniornis (originally dated to the Late Cretaceous); found at Ivo, Denmark. Ornithurae Scaniornithidae L. Cret. Eur. [nomen dubium]

Pasquiaornis Tokaryk, Cumbaa & Storer 1997 "Pasquia Hills bird"

PAS-kwee-a-OR-nis (Pasquia + Gr. ornis "bird") (m) named for the Pasquia Hills region of central Saskatchewan, Canada, where the specimens were found; a flightless diving bird more primitive that Baptornis, with a humerus and a femur that show resemblances to those of flying birds. Ornithurae Hesperornithiformes Baptornithidae L. Cret. NA.

Patagonykus Novas 1996 "Patagonian claw"

pat-a-GON-i-kus (Patagonia, region of southern Argentina (from Spanish patagon "big-foot," a name given the local Indians) + (Mon)onykus "one claw") (m) named to indicate a Mononykus-like animal found in the Patagonia region of Argentina. Classified by the author as a basal bird in the family Alvarezsauridae along with Mononykus and Alvarezsaurus, Patagonykus is considered a small theropod dinosaur by some other researchers. (?) Alvarezsauridae L. Cret. SA. [dino-bird]

Patagopteryx Alvarenga & Bonaparte 1992 "Patagonian wing"

pat-a-GOP-te-riks (Patagonia, region of southern Argentina, from Spanish patagon "big-foot," a name given the local Indians + Gr. pteryx "wing") (f) named for Patagonia, Argentina, where the fossils were found; a hen-sized flightless terrestrial bird, with a unique fused quadrate-pterygoid complex in its skull. Patagopterygidae L. Cret. SA.

Pelagornis Seeley 1866 "sea bird"

pel-a-GOR-nis (Gr. pelagos "sea" + Gr. ornis "bird") (m) for remains of a sea bird from the Cambridge Greensand marine deposits. (Preoccupied by Pelagornis Lartet 1857. See Enaliornis.) [nomen nudum]

Piksi Varricchio 2002 "big bird"

PIK-see (Blackfoot piksi "bird bird, chicken")* (m) named to indicate a fossil bird found on lands belonging to the Blackfeet Nation in Montana, interpreted "as a heavy-bodied ground bird with similarities to the Galliformes." Piksi is a medium-size bird, roughly the size of a modern ring-necked pheasant (Phasianus colchicus), known from a partial right humerus, ulna, and radius (Holotype: MOR 1113 (Museum of the Rockies, Bozeman, Montana)), found in the Late Cretaceous (Campanian) Two Medicine Formation, Glacier County, Montana. It differs from other known birds in having a large dorsal epicondyle on the humerus and large dorsal cotyla on the ulna; the ulna shaft appears to lack quill knobs. Piksi likely represents a heavy-bodied, ground-feeding bird that lived in an inland, relatively dry floodplain environment; it could only make relatively short strong flights based on wing features shared with modern tinamous and pheasants, birds that have limited flying abilities.

Type Species: Piksi barbarulna [bahr-bahr-OOL-nuh] Varricchio 2002: "strange-elbow" (from Lat. barabarus "strange" and Lat. ulna "elbow"), referring to the autapomorphic aspects of the new species (ulna with large dorsal cotyla twice as broad as ventral cotyla and with an elongate outline). Ornithothoraces i.s. Late Cretaceous (Campanian) NA. [added 6-2002]

Platanavis Nessov 1992 "sycamore bird"

pla-TAN-a-vis (Platanus Latin genus name for sycamore tree + Lat. avis "bird")* (f) alluding to the botanical genus Platanus (sycamore tree), dominant form of arboreal plant in the region of Central Kizylkum, Uzbekistan where the fossil was found (Dzhyrakuduk). i.s. L. Cret. CAs. (Uzbekistan)

Plegadornis Wetmore 1962 "ibis bird"

pleg-a-DOR-nis (Plegadis, Latin genus name for the ibis (from Gr. plegas "little sickle" for its bill) + Gr. ornis "bird") (m) originally identified as a "storklike bird," based on the shape of the humerus, and classified with ibises in the Threskiornithidae. (Preoccupied by Plegadornis Brehm 1855. See Angelinornis.) [= Ichthyornis]

Potamornis Elazanowski, Paul & Stidham 2001 "river bird"

pot-a-MOR-nis (Gr. potamos "river, stream" + Gr. ornis "bird") (m) probably alluding to Buck Creek, a tributary of Lance Creek, near the type locality in Wyoming, and to the near-shore stream environment where it may have lived. Potamornis is known from a quadrate bone (Holotype: UCMP 73103 (University of California Museum of Paleontology, Berkeley), apparently belonging to a small hesperornithiformid toothed bird; from the Late Cretaceous (Maastrichtian) Lance Formation in Niobrara County, Wyoming. Potamornis lived in a subtropical shoreline coastal environment during the final retreat of the inland sea that covered parts of western North America, and may have been a flightless, foot-propelled diver--the lack of pneumaticity in the bone suggests diving habits. Without more fossil material, however, the behavior and ecology of Potamornis remain unclear and it may have been different from typical hesperornithids.

Type Species: Potamornis skutchi [SKUH-chie] Elazanowski, Paul & Stidham 2001: in honor of Dr. Alexander F. Skutch, an eminent ornithologist, in recognition of his respect for birds' lives. Odontognathae ?Hesperornithiformes Late Cretaceous (Maastrichtian) NA [added 3-2001]

Protarchaeopteryx Ji Q. & Ji S. 1997 "first ancient-wing"

PROH-tahr-kee-OP-ter-iks (Gr. protos "first" + Archaeopteryx (Gr. arkhaios "ancient" + Gr. pteryx "wing")) (f) named to indicate a turkey-sized (3 ft. long) birdlike feathered animal similar to Archaeopteryx, but flightless and more primitive, originally described as coming between Archaeopteryx and the "feathered" dinosaur Sinosauropteryx in evolutionary terms. The fossils came from the famous lake bed deposits in the Liaoning region of northeastern China, in the Jiulongsong Member of the Chaomidianzi Formation, Jehol Group. Ji Qiang, P. Currie, M. Norell and Ji Shu-An redescribed the type specimen (Nature; 6-25-98) and concluded that Protarchaeopteryx was a maniraptoran theropod with feathers rather than a secondarily flightless primitive bird. Protarchaeopteryx resembles Caudipteryx in many features: the skull is relatively short; the hindlimbs are proportionately quite long and robust, suggesting a flightless running animal; the metatarsals (ankle bones) are not fused and resemble those of other theropods rather than birds; large symmetrical feathers form a fan at the end of the tail and bristle or down-like feathers covered most of the body; plumaceous feathers are preserved along the femurs. However, Protarchaeopteryx has serrated teeth in both the upper and lower jaws, with large straight premaxillary teeth, and short bulbous maxillary and dentary teeth; the forelimbs and the manus are longer than in Caudipteryx and all other long-armed coelurosaurs, but large arm feathers (found on the forelimbs and manus of the Caudipteryx specimens) are not preserved, and possibly may not have been present. The tail is longer than in Caudipteryx but has fewer than 28 vertebrae. No gizzard with gastroliths was found, indicating Protarchaeopteryx may have had a meat-eating diet (Caudipteryx had small gizzard stones and may have been primarily a plant-eater.)

Although Protarchaeopteryx is now classified as a dinosaur intermediate between typical ground-living maniraptoran coelurosaurs and primitive birds, a few researchers (some supporting a non-dinosaur origin for birds) suggest it is instead a secondarily flightless animal descended from a primitive Archaeopteryx-like flying form. The date for the fossils is still in dispute, but is either very Late Jurassic or very Early Cretaceous. Type species: Protarchaeopteryx robusta [roh-BUST-a] "robust, strong" for its large hindlimbs. Theropoda Coelurosauria Maniraptora E. Cret. China [dino-bird]

Protoavis Chatterjee 1986 "first bird"

PROH-to-AY-vis (Gr. protos "first" + Lat. avis "bird") (f) named to indicate its proposed status as a primitive ancestral bird, an identification disputed by many authorities. The occurrence of a bird in the Late Triassic would challenge the widely accepted theory that birds first appeared in the Late Jurassic. Although Chatterjee has described features of the vertebrae and skull as birdlike and provided a detailed reconstruction of the genus, the type material is fragmentary and no articulated specimens have been found to date. Most other researchers consider the currently available specimens too problematical to demonstrate that Protoavis was a bird, or even to establish that the fossils belong to a single kind of animal. It has been suggested instead that the material attributed to Protoavis comes from a number of Late Triassic reptiles, likely including one or more small theropod dinosaurs. John Ostrom (1991, 1996) has been a particularly sharp critic of Protoavis, and has found major shortcomings in the published descriptions. In the opinion of Luis Chiappe (1995), Protoavis "should not be considered relevant to avian evolution" until better specimens and articulated material are found, a position Ostrom endorses. Chatterjee (1995) has announced the discovery of additional well-preserved but disarticulated postcranial material that he attributes to Protoavis, but whether his on-going research will settle the controversy remains to be seen. Based on the published descriptions, the name Protoavis meets the basic requirements of nomenclature: it has a designated type specimen and a diagnosis that purports to define the taxon. The name is not preoccupied and so can be considered valid, although the taxonomic status of the genus is in dispute. Other researchers may consider the type material inadequate to define the genus for purposes of taxonomy and may question Chatterjee's interpretations of the fossils. Protoaviformes Protoavidae L. Trias. NA. [dino-bird]

Protopteryx Zhang & Zhou 2000 "primitive feather"

proh-TOP-te-riks (Gr. protos "first" + Gr. pteryx "wing," "feather") (f) referring to the apparently primitive nature of the long tail feathers, resembling unbranched scales. Protopteryx is a starling size enantiornithine bird known from a nearly complete skeleton with preserved feather impressions (slab and counterslab) (Holotype: IVPP V 11665 (Institute of Vertebrate Paleontology and Paleoanthropology)) and a second similar specimen, both from the Early Cretaceous (?Barremian) Yixian Formation Fengning County, Hebei Province, northern China. The skeletons show a mix of primitive and advanced features. The skull has a large orbit, pointed snout, with teeth in the upper and lower jaws; the neck has 7-8 cervical vertebrae; trunk has 12 dorsal vertebrae; synsacrum has 7 vertebrae; tail has 6-7 free caudal vertebrae. Unlike in other enantiornithines, there is a procoracoid process and a lateral process of the coracoid--features found in modern birds and associated with well-developed flight ability. The manus is longer than the forearm, with an unfused carpometacarpus and unreduced alular digit; the humerus and ulna are about of equal length. The femur is robust; the tibiotarsus is straight, unfused, and 120% the length of the femur. The toe claws are long and curved. The tail has a pygostyle. Three types of feathers are said to be present: flight feathers, down feathers, and a pair of "primitive, scalelike, central tail feathers." The authors consider the scalelike tail feathers (which appear to be unified without branching) a possible clue to the origin of feathers in birds, representing a transition for reptilian scales to complex feathers. Alternatively, the tail feathers in Protopteryx may be a secondary specialization, derived from ancestral normal feathers.

Type Species: Protopteryx fengningensis [fuhng-ning-EN-sis] Zhang & Zhou 2000: "from Fengning County" in Hebei Province, China. Enantiornithes Early Cretaceous (?Barremian) China [added 3-2001]

Rahona Forster, Sampson, Chiappe & Krause 1998 "menace from the clouds"

RAH-hoo-nah (Malagasy rahona "cloud, menace") (f) named to indicate a primitive bird from Madagascar with a large, hyperextensible "slashing" claw on the second digit of its feet as in dromaeosaurid dinosaurs. (Generic name preoccupied by an insect (Rahona Griveaud 1975); See Rahonavis]

Rahonavis Forster, Sampson, Chiappe & Krause 1998 "menace-from the-clouds bird"

rah-HOON-a-vis or rah-hoo-NAY-vis (Malagasy rahona "cloud, menace" + Lat. avis "bird") (f) named to indicate a primitive bird from Madagascar with a large, hyperextensible "slashing" claw on the second digit of its feet as in dromaeosaurid dinosaurs. (New name to replace preoccupied Rahona Forster, Sampson, Chiappe & Krause 1998). Skeletal material consists of a pelvis, femur, foot bones and digits, sacrum, dorsal vertebrae and part of the tail, associated with bones of a long forelimb bearing what appear to be quill knobs. The dorsal vertebrae have hyposphene-hypantra articulations, as in theropod dinosaurs. The pubis is vertical, not slanted back as in dromaeosaurids and birds, but has a hypopubic cup at the end, a feature associated with musculature that aids birds in ventilating their lungs. The metatarsal bones of the feet are not fused. These anatomical features, along with the large "slashing" claw on the foot, are very similar to dromaeosaurid dinosaurs. However, the apparent quill knobs on the long forelimbs indicate the animal had feathered wings and could fly. (Some researchers have questioned the association of the "wings" with the rest of the partial dinosaur-like skeleton. They suggest Rahonavis was a small, ground-living theropod dinosaur with a slashing claw, and that the wing bones may come instead from the partial remains of a more advanced type of bird (Vorona) found at the same fossil site, but with its wing bones missing.) Since the Madagascar site dates to the Late Cretaceous, Rahonavis as currently defined represents a very primitive "relict" bird with a long bony tail, similar to Archaeopteryx and dromaeosaurid dinosaurs. It was slightly larger than Archaeopteryx (about the size of a large modern crow). Type species: Rahonavis ostromi "for John Ostrom," the American vertebrate paleontologist who showed the similarities between Archaeopteryx and Deinonychus, and revived the theory that birds evolved from dinosaurs. Aves (?Theropoda) L. Cret. (?Campanian) Madagascar [dino-bird]

Sapeornis Zhou & Zhang 2002 "SAPE bird"

say-pee-OR-nis (SAPE (acronym for Society of Avian Paleontology and Evolution) + Gr. ornis "bird") (m) named in honor of the SAPE (Society of Avian Paleontology and Evolution), which held its fifth conference in China in June 2000, shortly before the holotype specimen was collected. Sapeornis is the largest known Early Cretaceous bird (around the size of a large gull; or roughly a third larger than Archaeopteryx in most details except the tail and hindlimbs, and about twice as large as the dromaeosaur Microraptor), based on a nearly complete postcranial skeleton (Holotype V 12698 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China)), including the vertebral column, pectoral and pelvic girdles, partially articulated forelimbs, left hindlimb, and fused pygostyle; found in the Early Cretaceous (Barremian?) Jiufotang Formation, at Shangheshou, Chaoyang City, Liaoning Province, northeast China. The forelimbs are proportionately extremely long while the hindlimbs are relatively short (forelimb to hindlimb ratio 1.55), features that along with the well-developed deltoid crest of the humerus, proximally fused metacarpals, and short pygostyle (at least 4 fused vertebrae), indicate powerful soaring capability, suggesting Sapeornis was a long-winged soaring shorebird. Elongated heterocoelous cervical vertebrae gave it a flexible neck. Sapeornis is the most basal (primitive) bird known except for Archaeopteryx, and shows a mosaic of advanced and primitive features compared to other Early Cretaceous birds: like Archaeopteryx and dromaeosaurs, it has a short, nonstrut-like coracoid, well developed manual digit I, fibula reaching the distal end of the tarsal joint, a free tarsal and a 5th metatarsal: it is more advanced than Archaeopteryx in having heterocoelous neck vertebrae, short tail with a pygostyle, fused carpo-metacarpus, pubic symphysis 1/3 length of the pubis, and ulna longer than radius. The discovery of a primitive bird well-adapted to soaring suggests that the ecological diversity of Early Cretaceous birds was greater than previously thought.

Type species: Sapeornis chaoyangensis [chow-yahng-EN-sis] Zhou & Zhang 2001: "from Chaoyang" for the type locality at Chaoyang.Aves (order indeterminate) Early Cretaceous (Barremian?) China [added 6-2002]

Saururae Haeckel 1866 "lizard tails"

saw-ROOR-ee (Gr. sauros "lizard" + Gr. oura "tail") (f) Proposed for Archaeopteryx, a bird with a long reptilian tail, to distinguish it from the Ornithurae, with shortened tails. Now used with the spelling "Sauriurae" by some researchers (Fedducia) for a group including Archaeopteryx and the Enantiornithes. [Taxon]

Sazavis Nessov 1989 "clay bird"

SAZ-a-vis (Kazakh saz "clay" + Lat. avis "bird")* (f) named for a clay depression near where the fossil was found in the central Kizylkum Desert at Dzharakuduk, Uzbekistan. Enantiornithes Alexornithidae L. Cret. CAs.

Shuvuuia Chiappe, Norell & Clark 1998 "bird"

shu-VOO-ee-a (Mong. shuvuu "bird" + -ia) (f) named to indicate a Mononykus-like animal from Mongolia (South Gobi Aimak (Djadokhta Formation)), notable for many birdlike features in its skull, including the ability to bend the snout upward independently of the braincase (prokinesis). The skull is about 8 cm. (3.2 in.) long, and delicately built with large orbits and tiny, unserrated teeth in the front part of the upper jaws and along a continuous groove in the dentary bone of the lower jaw. The material consists of two extremely well preserved skulls, with vertebrae, limbs, and other post-cranial bones originally attributed to Mononykus. Shuvuuia is distinguished from Mononykus by the shape of its humerus and tibia, as well as its neck vertebrae and skull, but was similar in size (about the size of a wild turkey). To date, Shuvuuia provides the best skull material from any alvarezsaurid, and, according to the authors' cladistic analysis, confirms the identification of alvarezsaurids as a family of primitive flightless birds rather than as a group of non-avian theropod dinosaurs. Type species: Shuvuuia deserti [dee-SER-tie] "of the desert," referring to the semi-arid environment in which the Djadokhta Formation (?Campanian) was deposited. Theropoda (Aves) Alvarezsauridae Mononykinae L. Cret. Mongolia [dino-bird]

Sinornis Sereno & Rao 1992 "Chinese bird"

sie-NOR-nis (New Lat. sino- (Gr. Sinai (Lat. Sinae) an oriential race) "Chinese" + Gr. ornis "bird") (m) named for China, where the sparrow-size specimen was found in a lake bed deposit in Liaoning Province (northeastern China). (?) Enantiornithes E. Cret. China

Songlingornis Hou 1997 "Songling (Mountains) bird"

soong-ling-OR-nis (Chin. Songling + Gr. ornis "bird") (m) named for the Songling Mountains northeast of the Chaoyang region of Liaoning Province; an ornithurine toothed bird about the size of a sparrow, known from an incomplete skeleton and parts of a skull (Holotype: IVPP V. 10913) from the Jiufotang Formation at Boluochi, Chaoyang County, Liaoning Province, northeastern China. Dense arrangement of teeth along the length of the upper and lower jaws; coracoid well developed; furcula process narrow as in modern waterbirds; sternum large, with carina, fenestrae and lateral processes; ulna 2.8 cm long, radius 2.1 cm long.

Type Species: Songlingornis linghensis [ling-HEN-sis] Hou 1997, referring to the Ling River (consisting of the Daling He "Big Ling River" and the Xiaoling He "Little Ling River"), running in a southwest to northeast direction near the region of Liaoxi (western Liaoning Province) in which the fossil was found in northern China. Ornithurae Chaoyangiformes Songlingornithidae Early Cretaceous (?Barremian) China [added 2/2000]

Soroavisaurus Chiappe 1993 "sister-avisaur"

SOR-o-AY-vi-SAWR-us (Lat. soror "sister" + Avisaurus)* (m) referring to the sister-group relationship inferred for Avisaurus and Soroavisaurus, according to a cladistic analysis. Enantiornithes Avisauridae L. Cret. SA.

Telmatornis Marsh 1870 "marsh bird"

TEL-ma-TOR-nis (Gr. telmat- (telma) "marsh, swamp" + Gr. ornis "bird") (m) named to indicate a form "probably related to [the Rail family]," found in the Cretaceous Greensand of New Jersey. Ornithurae ?Charadriiformes Graculavidae L. Cret. NA.

Torotix Brodkob 1963 "flamingo"

TOR-o-tiks (Gr. torotix, imitation of a bird's cry, attributed to a flamingo by the ancient Greek comic playwright Aristophanes)* (f) originally identified as the femur bone of a flamingo; its true classification is disputed. Ornithurae Torotigidae. L. Cret. NA. [nomen dubium]

Tytthostonyx Olson & Parris 1987 "little spur"

ti-THOS-to-niks (Gr. tytthos "little" + Gr. stonyx "small point") (m) name "refers to the small, presumably rudimentary, ectepicondylar spur" on the humerus. Ornithurae ?Procellariiformes Tytthostonychidae L. Cret. NA.

Volgavis Nessov & Jarkov 1989 "Volga River bird"

VOL-ga-vis (Volga + Lat. avis "bird") (f) named for the Volga River Basin, Volograd District, southern Russia, where the fossil was found; with a hook bill oriented downward, in toothless jaws that show some similarities to primitive charadriiforms and the Fregatidae (frigate birds). Ornithurae Charadriiformes i.s. L. Cret. EEur.

Vorona Forster, Chiappe, Krause & Sampson 1996 "bird"

voo-ROO-na (Malagasy vorona "bird")* (f) named to indicate a fossil bird from Madagascar; for well preserved leg bones found near the village of Berivota, northwest Madagascar. Ornithothoraces L. Cret. Madagascar

Yandangornis Cai & Zhao 1999 "Yandang bird"

yen-dahng-OR-nis (Chin. Yangang + Gr. ornis "bird") (m) named for the Yandang Mountains, where the fossil was collected at the foot of the northeast slope, in Zhejiang Province, eastern China. Yandangornis is known from a nearly complete skeleton (Holotype: M1326 (Zhejiang Museum of Natural History)) found in a stone pit near Aoli Village, Shangpan Town, Linhai City, Zhejiang Province, China, and dated to the early Late Cretaceous (?Santonian) from the Tangshang Group horizon. The remains indicate a ground-living bird (or possibly a small, very birdlike theropod dinosaur) about the size of Archaeopteryx (crow-size), with a similar long, primitive bony tail but differing most in having a lightly built skull with toothless jaws. The total length is about 588 mm (around 2 ft): 283 mm from skull to pubis; at least 305 mm for tail. The skull is about 50 mm long, fairly wide and short; the mandible is narrow and thin. The jaws are toothless and had a horny beak in life. The neck is about 80 mm long, with biconcave vertebrae without cervical ribs; the anterior vertebrae are shorter than the posterior ones. The forelimbs are large but incompletely preserved, with a humerus with no pneumatic fossa or pneumatic foramen; the bones in the manus are separate and two digits are free. The sternum is large (50 mm long), has posterior and lateral processes but lacks a keel. Abdominal ribs are present. The pubis is inclined backwards, with no anterior process. The hindlimbs are well-developed: the femur (106 mm long) is 3/4th the length of the tibiotarsus (132 mm long) but about 1/3 longer than the relatively short and wide tarsometatarsus (70 mm long). The toes and hallux are short, with small, uncurved claws. The tail is long (over half the total length of the body), consisting of more than 20 caudals (19 preserved) without haemal arches. Apparently no traces of feathers were preserved with the fossil.

The short hallux and uncurved claws show Yandangornis was not built to perch in trees; the heavy bony tail may have limited or prevented any possible flying ability. The authors identify Yandangornis as a flightless terrestrial bird showing a peculiar mix of advanced and primitive features (a beaked, Confuciusornis-like toothless skull but Archaeopteryx-like long, bony tail). Some researchers who have seen the specimen consider the animal an unusual small theropod dinosaur, but the authors cite the wide sternum, details of the hindlimbs, and lack of haemal arches on the tail as reasons for classifying Yandangornis as a bird.

Type Species: Yandangornis longicaudus [lon-ji-KAW-duhs] Cai & Zhao 1999: "long-tailed" Yandangornithiformes Yandangornithidae Late Cretaceous (Santonian) China [dino-bird][added 7/2000]

Yanornis Zhou & Zhang 2001 "Yan (Dynasty) bird"

yen-OR-nis (Chin. Yan (name of a dynasty) + Gr. ornis "bird") (m) name "derived from the Chinese spelling of the ancient Chinese Yan Dynasty [usually called Former Yan Dynasty (337-370 AD)], the type locality Chaoyang City was its capital" in modern Liaoning Province in northeastern China. Yanornis is a crow-size ornithurine bird known from a nearly complete skeleton without feather impressions (Holotype: V12558 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), found in the Early Cretaceous (Barremian?) Jiufotang Formation at Chaoyang City, Yixian County in Liaoning Province, northeast China. Although the sternum, coracoid and scapula closely resemble those of modern birds and indicate well developed flying abilities, Yanornis retains a number of primitive features, including robust conical teeth in the jaws, claws on the first two digits of the manus, and a long pubic symphysis. The neck has at least 10 heterocoelous neck vertebrae; preserved dorsal vertebrae have long and deep pleurocoels; synsacrum has 9 sacral vertebrae; pygostyle is short. The length from tip of jaws to end of pygostyle can be estimated at around 38-40 cm (15-16 in). The slender, backward-directed pubis has a pubic symphysis about 30% the total length. Yanornis's long, rather narrow toothed jaws and relatively long neck indicate that it preyed on fish and other aquatic animals; its long toes with short claws and reduced hallux suggest it spent much of its time walking along the shore rather than perching in trees.

Type Species: Yanornis martini [MAHR-ti-nie] Zhou & Zhang 2001: for "Larry D. Martin [American vertebrate paleontologist] for his contributions to the study of Mesozoic birds." Ornithurae Yanornithiformes Yanornithidae Early Cretaceous (Barremian?) China [added 8-2001]

Yixianornis Zhou & Zhang 2001 "Yixian bird"

yee-shyen-OR-nis (Chin. Yixian (County) + Gr. ornis "bird") (m) named for the locality Yixian County in Liaoning Province, where the fossils were found. Yixianornis is an ornithurine bird roughly the size of a blue-jay, known from a nearly complete specimen with feather impressions (Holotype: V12631 (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing)), found in the Early Cretaceous (Barremian?) Jiufotang Formation at Qianyang in Yixian County, Liaoning Province, northeast China. Yixianornis has strong flying capabilities indicated by its long wings and by a sternum, coracoid and scapula similar in form to modern birds, but it also retains primitive features such as small teeth in the jaws, claws on the first two wing-fingers, a pubic symphysis, and gastralia--the first gastralia documented for an ornithurine bird. The skull is relatively broad and short, with a length to width ratio of 1.5. The neck has at least 9 heterocoelous vertebrae. The tail has 5 short free caudals and a short pygostyle; preserved tail feathers are short. The length from tip of jaws to end of pygostyle was probably around 20-22 cm (8-9 in). The hindlimb and toe bones are relatively long and slender, with only a small hallux, suggesting it spent time on the ground or along the shore rather than perched in trees.

Type Species: Yixianornis grabaui [GRAB-ow-ie] Zhou & Zhang 2001: "for the American geologist Amadeus William Grabau [1870-1946], a pioneering geologist in Liaoning, for his contribution to the study of Jehol Biota." Ornithurae Chaoyangiornithiformes Early Cretaceous (Barremian?)China [added 8-2001]

Yungavolucris Chiappe 1993 "Yunga (Argentina) bird"

YOONG-ga-vo-LOO-kris (Yunga + Lat. volucris "bird")* (f) referring to Yunga, the phytogeographic region in which El Brete type location is found in Salta Province, northwestern Argentina. Enantiornithes i.s. L. Cret. AS.

Zhyraornis Nessov 1984 "Dzhyrakuduk (Uzbekistan) bird"

JAH-ray-OR-nis (Zhyra [= Dzhyra] + Gr. ornis "bird") (m) named for the Dzhyrakuduk waterwell near where the type fossil was found, Tamdy district, Central Kyzylkum, Uzbekistan. Ornithurae Zhyraornithidae L. Cret. Cas.